The effects of Mid-Holocene foragers on the European oyster in Denmark.

Oysters (Ostreidae) play a pivotal role in the health and productivity of marine ecosystems. Their unique ability to filter water, provide habitat, and contribute to nutrient cycling has remained underused in many parts of Europe following the destruction of vast oyster beds in the 19th and 20th centuries. The burgeoning field of oyster restoration for aquaculture has recognized the potential of these bivalves in promoting ecosystem resilience and enhancing biodiversity. Restoring oysters to previous levels requires the establishment of ecological baselines that ideally take into account the long-term changes of animal behavior as well as the surrounding environment prior to significant human intervention, an extremely challenging task. Archaeological shell middens are invaluable baseline archives and provide exclusive insights into past ecosystems. Here, we use demographic information from over 2,000 analyzed European oyster (Ostrea edulis) shells dating from ~5,660 to 2,600 cal BCE (calibrated years BCE), the largest archaeological growth rate dataset of mollusks yet. Through the analysis of size as well as ontogenetic age, we decouple anthropogenic from environmental impacts throughout Denmark. Our data show definitive influence of oyster size-age structure through human harvesting during the Mid-Holocene, with older oysters in the Mesolithic (mean: 4.9 y) than the Neolithic (mean: 3.7 y), irrespective of changes in growth rate. Furthermore, we present the metrics for long-term sustainable harvesting of oysters across environmental and socioeconomic transitions, providing demographic targets for current oyster restoration projects and valuable context in mitigating the impact of modern climatic change.

The impact of farming on prehistoric culinary practices throughout Northern Europe.

To investigate changes in culinary practices associated with the arrival of farming, we analysed the organic residues of over 1,000 pottery vessels from hunter-gatherer-fisher and early agricultural sites across Northern Europe from the Lower Rhine Basin to the Northeastern Baltic. Here, pottery was widely used by hunter-gatherer-fishers prior to the introduction of domesticated animals and plants. Overall, there was surprising continuity in the way that hunter-gatherer-fishers and farmers used pottery. Both aquatic products and wild plants remained prevalent, a pattern repeated consistently across the study area. We argue that the rapid adaptation of farming communities to exploit coastal and lagoonal resources facilitated their northerly expansion, and in some cases, hunting, gathering, and fishing became the most dominant subsistence strategy. Nevertheless, dairy products frequently appear in pottery associated with the earliest farming groups often mixed with wild plants and fish. Interestingly, we also find compelling evidence of dairy products in hunter-gatherer-fisher Ertebølle pottery, which predates the arrival of domesticated animals. We propose that Ertebølle hunter-gatherer-fishers frequently acquired dairy products through exchange with adjacent farming communities prior to the transition. The continuity observed in pottery use across the transition to farming contrasts with the analysis of human remains which shows substantial demographic change through ancient DNA and, in some cases, a reduction in marine consumption through stable isotope analysis. We postulate that farmers acquired the knowledge and skills they needed to succeed from local hunter-gatherer-fishers but without substantial admixture.

Deep antiquity of seagrasses supporting European eel fisheries in the western Baltic.

Protecting ocean habitats is critical for international efforts to mitigate climate impacts and ensure food security, but the ecological data upon which policy makers base conservation and restoration targets often reflect ecosystems that have already been deeply impacted by anthropogenic change. The archaeological record is a biomolecular archive offering a temporal scope that cannot be gathered from historical records or contemporary fieldwork. Insights from biogeochemical and osteometric analyses of fish bones, combined with context from contemporary field studies, show how prehistoric fisheries in the western Baltic relied on seagrass meadows. European eels (Anguilla anguilla) harvested by Mesolithic and Neolithic peoples over millennia showed a strong fidelity for eelgrass foraging habitats, an ecological relationship that remains largely overlooked today, demonstrating the value of protecting these habitats. These data open new windows onto ecosystem- and species-level behaviours, highlighting the need for wider incorporation of archaeological data in strategies for protecting our oceans.

The impact of environmental change on the use of early pottery by East Asian hunter-gatherers.

The invention of pottery was a fundamental technological advancement with far-reaching economic and cultural consequences. Pottery containers first emerged in East Asia during the Late Pleistocene in a wide range of environmental settings, but became particularly prominent and much more widely dispersed after climatic warming at the start of the Holocene. Some archaeologists argue that this increasing usage was driven by environmental factors, as warmer climates would have generated a wider range of terrestrial plant and animal resources that required processing in pottery. However, this hypothesis has never been directly tested. Here, in one of the largest studies of its kind, we conducted organic residue analysis of >800 pottery vessels selected from 46 Late Pleistocene and Early Holocene sites located across the Japanese archipelago to identify their contents. Our results demonstrate that pottery had a strong association with the processing of aquatic resources, irrespective of the ecological setting. Contrary to expectations, this association remained stable even after the onset of Holocene warming, including in more southerly areas, where expanding forests provided new opportunities for hunting and gathering. Nevertheless, the results indicate that a broader array of aquatic resources was processed in pottery after the start of the Holocene. We suggest this marks a significant change in the role of pottery of hunter-gatherers, corresponding to an increased volume of production, greater variation in forms and sizes, the rise of intensified fishing, the onset of shellfish exploitation, and reduced residential mobility.

Archaeological bone lipids as palaeodietary markers.

RATIONALE: Stable isotope analysis of archaeological and fossil bone samples can provide important insights into past environments, ecologies and diets. Previous studies have focused on stable carbon and nitrogen isotopes in bone collagen, or carbon isotopes in bone mineral (bioapatite). Carbon isotope analysis of lipids from archaeological bone has received much less attention, partly due to the lack of suitable methodologies allowing sufficient recovery of compounds for structural and isotopic characterisation. Here we show that lipids can be easily and reliably recovered from archaeological bone using a modified protocol, and that these provide complementary dietary information to other bone components. METHODS: Human and animal bones were obtained from a variety of archaeological contexts. Lipids were sequentially extracted using solvent extraction (dichloromethane/methanol), followed by acidified methanol extraction (methanol/H2SO4). The lipids were then analysed by gas chromatography/mass spectrometry (GC/MS) and gas chromatography/combustion/isotope ratio mass spectrometry (GC/C/IRMS). RESULTS: Appreciable amounts of endogenous lipid were recovered from archaeological bone. Importantly, a comparison between compound-specific and bulk collagen isotopic data shows that archaeological bone lipids reflect dietary input and can be used to distinguish between marine and terrestrial consumers, as well as between C3 and C4 plant consumers. Furthermore, the presence of essential fatty acids directly incorporated from diet to bone may provide additional palaeodietary information. CONCLUSIONS: Our findings suggest that archaeological bone lipids are a hitherto untapped resource of dietary information that offer additional insights to those gained from other isotopic analyses of bone.

Carbon, nitrogen and sulphur isotope data of archaeological fish and mammal bone collagen from Lithuania.

Until relatively recently, stable sulphur isotope analysis of bone collagen was seldom undertaken in bioarchaeological research. With increasing frequency, its application has proven useful in reconstructing palaeodiets and palaeoecologies, as well as identifying potential migration and mobility patterns. Here, sulphur (δ34S) isotope analysis, together with carbon (δ13C) and nitrogen (δ15N), was performed on six fish and 34 mammal bone collagen samples from 14 prehistoric sites in Lithuania dating from the Late Mesolithic (ca. 7000-5000 cal BC) to the Late Bronze Age (ca. 1100-500 cal BC). We present the first δ34S data from Lithuania, including coupled δ13C and δ15N data, offering a crucial dataset for future research to explore spatial and temporal variability in the region and beyond.

Detection of dairy products from multiple taxa in Late Neolithic pottery from Poland: an integrated biomolecular approach.

The detection of dairy processing is pivotal to our understanding of ancient subsistence strategies. This culinary process is linked to key arguments surrounding the evolution of lactase persistence in prehistory. Despite extensive evidence indicating the presence of dairy products in ceramics in the European Neolithic, questions remain about the nature and extent of milk (and lactose) processing and consumption. In order to investigate past patterns of dairy processing, here we analyse ancient proteins identified from Late Neolithic Funnel Beaker ceramics, scrutinizing the principle that curd and whey proteins partition during the production of dairy foods from milk. Our results indicate the presence of casein-rich dairy products in these vessels suggesting the creation of curd-enriched products from raw milk. Moreover, this analysis reveals the use of multiple species for their dairy products in the Late Neolithic, adding to a growing body of evidence for the period. Alongside palaeoproteomic analysis, we applied well-established lipid residue analysis. Differential interpretations between these two approaches show that palaeoproteomics is especially useful where the effects from isotope mixing may underestimate the frequency of dairy products in archaeological ceramics, highlighting the potential utility of a multi-stranded approach to understand life histories of vessel use.

Parallel worlds and mixed economies: multi-proxy analysis reveals complex subsistence systems at the dawn of early farming in the northeast Baltic.

The transition from foraging to farming was a key turning point in ancient socio-economies. Yet, the complexities and regional variations of this transformation are still poorly understood. This multi-proxy study provides a new understanding of the introduction and spread of early farming, challenging the notions of hierarchical economies. The most extensive biological and biomolecular dietary overview, combining zooarchaeological, archaeobotanical, dietary stable isotope and pottery lipid residue analyses is presented, to unravel the nature and extent of early farming in the 3rd millennium cal BCE in the northeast Baltic. Farming was introduced by incoming Corded Ware cultural groups (CWC), but some dietary segregation existed within these communities, with some having more access to domesticates, others incorporating more wild resources into their diet. The CWC groups coexisted in parallel with local hunter-fisher-gatherers (HFG) without any indication of the adoption of domesticates. There was no transition from foraging to farming in the 3rd millennium cal BCE in the NE Baltic. Instead, we see a complex system of parallel worlds with local HFGs continuing forager lifeways, and incoming farmers practising mixed economies, with the continuation of these subsistence strategies for at least a millennium after the first encounter with domesticated animals.

The transmission of pottery technology among prehistoric European hunter-gatherers.

Human history has been shaped by global dispersals of technologies, although understanding of what enabled these processes is limited. Here, we explore the behavioural mechanisms that led to the emergence of pottery among hunter-gatherer communities in Europe during the mid-Holocene. Through radiocarbon dating, we propose this dispersal occurred at a far faster rate than previously thought. Chemical characterization of organic residues shows that European hunter-gatherer pottery had a function structured around regional culinary practices rather than environmental factors. Analysis of the forms, decoration and technological choices suggests that knowledge of pottery spread through a process of cultural transmission. We demonstrate a correlation between the physical properties of pots and how they were used, reflecting social traditions inherited by successive generations of hunter-gatherers. Taken together the evidence supports kinship-driven, super-regional communication networks that existed long before other major innovations such as agriculture, writing, urbanism or metallurgy.

High-resolution dietary reconstruction of victims of the 79 CE Vesuvius eruption at Herculaneum by compound-specific isotope analysis.

The remains of those who perished at Herculaneum in 79 CE offer a unique opportunity to examine lifeways across an ancient community who lived and died together. Historical sources often allude to differential access to foodstuffs across Roman society but provide no direct or quantitative information. By determining the stable isotope values of amino acids from bone collagen and deploying Bayesian models that incorporate knowledge of protein synthesis, we were able to reconstruct the diets of 17 adults from Herculaneum with unprecedented resolution. Significant differences in the proportions of marine and terrestrial foods consumed were observed between males and females, implying that access to food was differentiated according to gender. The approach also provided dietary data of sufficient precision for comparison with assessments of food supply to modern populations, opening up the possibility of benchmarking ancient diets against contemporary settings where the consequences for health are better understood.

Latitudinal gradient in dairy production with the introduction of farming in Atlantic Europe.

The introduction of farming had far-reaching impacts on health, social structure and demography. Although the spread of domesticated plants and animals has been extensively tracked, it is unclear how these nascent economies developed within different environmental and cultural settings. Using molecular and isotopic analysis of lipids from pottery, here we investigate the foods prepared by the earliest farming communities of the European Atlantic seaboard. Surprisingly, we find an absence of aquatic foods, including in ceramics from coastal sites, except in the Western Baltic where this tradition continued from indigenous ceramic using hunter-gatherer-fishers. The frequency of dairy products in pottery increased as farming was progressively introduced along a northerly latitudinal gradient. This finding implies that early farming communities needed time to adapt their economic practices before expanding into more northerly areas. Latitudinal differences in the scale of dairy production might also have influenced the evolution of adult lactase persistence across Europe.

Correction to 'Organic residue analysis shows sub-regional patterns in the use of pottery by Northern European hunter-gatherers'.

[This corrects the article DOI: 10.1098/rsos.192016.].

Organic residue analysis shows sub-regional patterns in the use of pottery by Northern European hunter-gatherers.

The introduction of pottery vessels to Europe has long been seen as closely linked with the spread of agriculture and pastoralism from the Near East. The adoption of pottery technology by hunter-gatherers in Northern and Eastern Europe does not fit this paradigm, and its role within these communities is so far unresolved. To investigate the motivations for hunter-gatherer pottery use, here, we present the systematic analysis of the contents of 528 early vessels from the Baltic Sea region, mostly dating to the late 6th-5th millennium cal BC, using molecular and isotopic characterization techniques. The results demonstrate clear sub-regional trends in the use of ceramics by hunter-gatherers; aquatic resources in the Eastern Baltic, non-ruminant animal fats in the Southeastern Baltic, and a more variable use, including ruminant animal products, in the Western Baltic, potentially including dairy. We found surprisingly little evidence for the use of ceramics for non-culinary activities, such as the production of resins. We attribute the emergence of these sub-regional cuisines to the diffusion of new culinary ideas afforded by the adoption of pottery, e.g. cooking and combining foods, but culturally contextualized and influenced by traditional practices.

Diet, cuisine and consumption practices of the first farmers in the southeastern Baltic.

With the arrival of the Early Neolithic Globular Amphora and Corded Ware cultures into the southeastern Baltic, ca. 2900/2800-2400 cal BC, a new type of economy was introduced, animal husbandry. However, the degree to which this transformed the subsistence economy is unknown. Here, we conducted organic residue analyses of 64 ceramic vessels to identify their contents. The vessels were sampled from 10 Lithuanian archaeological sites dating across the Subneolithic-Neolithic transition to the Early Bronze Age (ca. 2900/2800-1300 cal BC). Our results demonstrate that regardless of location or vessel type, many ceramics were used to process aquatic resources. Against our expectations, this association continued even after marked economic change concurrent with the migration of pastoralists from central and southeastern Europe, as evidenced by recent ancient DNA analysis of human remains. Moreover, we observed dairy fats in pottery from all cultures of the Early Neolithic (i.e. Rzucewo, Globular Amphora and Corded Ware) but unlike other regions of Europe, it seems that these were incorporated into indigenous culinary practices. Furthermore, some vessels were used to process plant foods, and others may have been used for the production and/or storage of birch bark tar. However, evidence for domesticated plant processing, for example millet, was absent. We show that organic residue analysis of pottery provides a different picture of past consumption patterns compared to the stable isotope analysis of human remains from isolated burials where a clear dietary shift is evident.

Dental calculus and isotopes provide direct evidence of fish and plant consumption in Mesolithic Mediterranean.

In this contribution we dismantle the perceived role of marine resources and plant foods in the subsistence economy of Holocene foragers of the Central Mediterranean using a combination of dental calculus and stable isotope analyses. The discovery of fish scales and flesh fragments, starch granules and other plant and animal micro-debris in the dental calculus of a Mesolithic forager dated to the end of the 8th millenium BC and buried in the Vlakno Cave on Dugi Otok Island in the Croatian Archipelago demonstrates that marine resources were regularly consumed by the individual together with a variety of plant foods. Since previous stable isotope data in the Eastern Adriatic and the Mediterranean region emphasises that terrestrial-based resources contributed mainly to Mesolithic diets in the Mediterranean Basin, our results provide an alternative view of the dietary habits of Mesolithic foragers in the Mediterranean region based on a combination of novel methodologies and data.

Early Neolithic genomes from the eastern Fertile Crescent.

We sequenced Early Neolithic genomes from the Zagros region of Iran (eastern Fertile Crescent), where some of the earliest evidence for farming is found, and identify a previously uncharacterized population that is neither ancestral to the first European farmers nor has contributed substantially to the ancestry of modern Europeans. These people are estimated to have separated from Early Neolithic farmers in Anatolia some 46,000 to 77,000 years ago and show affinities to modern-day Pakistani and Afghan populations, but particularly to Iranian Zoroastrians. We conclude that multiple, genetically differentiated hunter-gatherer populations adopted farming in southwestern Asia, that components of pre-Neolithic population structure were preserved as farming spread into neighboring regions, and that the Zagros region was the cradle of eastward expansion.